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The Reproductive Biology of Eleutherodactylus Jasperi (Amphibia, Anura, Leptodactylidae), with Comments on the Evolution of Live-Bearing Systems

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The Reproductive Biology of Eleutherodactylus Jasperi (Amphibia, Anura, Leptodactylidae), with Comments on the Evolution of Live-Bearing Systems The Reproductive Biology of Eleutherodactylus jasperi (Amphibia, Anura, Leptodactylidae), with Comments on the Evolution of Live-Bearing Systems Marvalee H. Wake Journal of Herpetology, Vol. 12, No. 2. (Apr. 24, 1978), pp. 121-133. Stable URL: http://links.jstor.org/sici?sici=0022-1511%2819780424%2912%3A2%3C121%3ATRBOEJ%3E2.0.CO%3B2-F Journal of Herpetology is currently published by Society for the Study of Amphibians and Reptiles. Your use of the JSTOR archive indicates your acceptance of JSTOR's Terms and Conditions of Use, available at http://www.jstor.org/about/terms.html. JSTOR's Terms and Conditions of Use provides, in part, that unless you have obtained prior permission, you may not download an entire issue of a journal or multiple copies of articles, and you may use content in the JSTOR archive only for your personal, non-commercial use. Please contact the publisher regarding any further use of this work. Publisher contact information may be obtained at http://www.jstor.org/journals/ssar.html. Each copy of any part of a JSTOR transmission must contain the same copyright notice that appears on the screen or printed page of such transmission. The JSTOR Archive is a trusted digital repository providing for long-term preservation and access to leading academic journals and scholarly literature from around the world. The Archive is supported by libraries, scholarly societies, publishers, and foundations. It is an initiative of JSTOR, a not-for-profit organization with a mission to help the scholarly community take advantage of advances in technology. For more information regarding JSTOR, please contact [email protected]. http://www.jstor.org Sun Sep 9 15:05:52 2007 JOURNAL OF HERPETOLOGY Volume 12 24 April 1978 Number 2 The Reproductive Biology of Eleutherodactylus jasperi (Amphibia, Anura, Leptodactylidae), with Comments on the Evolution of Live-bearing Systems Marvalee H. Wake Department of Zoology and Museum of Vertebrate Zoology, University of California, Berkeley CA 94720, USA ABSTRACT-Eleutherodactylus jasperi females retain developing embryos in the oviducts. Three to five fully metamorphosed froglets are born approximately 33 days after the female was in amplexus. Internal fertilization is presumed. Adult morphology of the reproductive organs is not unique. The oviduct has two distinct regions. Clutch size is very small (1-6 ova). The development of the froglets is very similar to that described for several species of direct developing Eleutherodactylus. The froglets have a small egg tooth. The tail is thin, highly vascufarized, and overlies the dorsum or venter of the embryo. It may function as an organ for gaseous exchange with the oviducal capillaries. Unresorbed yolk is present in froglets even after birth. No other maternal nutrition is suspected. E. jasperi reproductive biology is compared with that of Nectophrynoides and with trends in reproductive biology within the genus Eleutherodactylus. INTRODUCTION Members of the genus Eleutherodactylus are terrestrial frogs, all of which are direct developers that lack a free-living larval stage. Most species for which the reproductive biology is known lay clutches of relatively few (10-104) large yolky eggs in protected sites on the ground, on leaves, or in bromeliads; fully metamorphosed froglets hatch from these eggs. However, one species of Eleutherodactylus, the recently described E. jasperi (Drewry and Jones, 1976), is ovoviviparous. The species lives in bromeliads in the mountains south of Cayey, Puerto Rico, at elevations above 700 m. Females of the species retain the developing eggs in a chamber formed of the fused portions of the oviducts, and fully metamorphosed froglets are subsequently born. This report describes the gross and microscopic morphology of the adult male and female reproductive systems, the anatomy of the later developmental stages of the froglets, and discusses the acquisition of ovoviviparity as a reproductive mode and its evolutionary significance. MATERIALS AND METHODS Nine adult specimens of E. jasperi were available for study. Eight (four males and four females; three of the latter carrying oviducal froglets) were preserved in formalin in the field and transferred to 70% ethanol. One specimen was preserved in the laboratory in Zenker's formol in order to better control potential shrinkage. In addition, seven newborn and thirteen oviducal froglets were available. Table 1 summarizes information about the specimens and their manner of study. 1978 JOURNAL OF HERPETOLOGY 12(2):121-133 (121) MARVALEE H. WAKE TABLE 1. Material of Eleutherodactylus jasperi Available for Study Reproductive organs and [Brackets associate members of clutches with the maternal female.] froglets were sectioned at 10 p and stained with hematoxylin- Manner of Study Specimens eosin or picro-ponceau. All -Snout-vent Length measurements were read with a Sex (mm) Dissection Histological Preparation dial caliper and most dissections d 17.4 x ~ ~kidney,ducts ~ ~ werei ~done , with the aid of a d 18.2 x Wild dissecting microscope. d 19.1 x Testis, kidney, ducts d 18.8 x Testis, kidney, fat body 9 19.4 x Ovary, kidney, ducts 9 20.5 x Ovary, kidney GROSS MORPHOLOGY OF 0 20.7 x Ovary, oviduct, cloaca THE FEMALE REPRO- 9 20.9 X 0 21.5 x Ovary, oviducts DUCTIVE TRACT Newborn 6m The ovaries.-The ovaries are the paired sacciform struc- Newborn 7.1 -20.7 9 Newborn 7.2 Entire tures typical of most frogs, but Newborn they do not fill the abdominal Newborn Entire cavity since an ovary contains a Oviducal maximum of six fully yolked Oviducal Entire large (to 3.3 mm diameter) Oviducal Two 'unmeasured eggs. The ovary walls contain Oviducal large melanophores, and the Oviducal Entire presence of eggs under the pig- Oviducal mented wall visible through the Oviducal skin of the venter is used in the Oviducal field to determine that ovula- Oviducal (with membrane) Entire Oviducal (with membrane) Entire tion has not occurred (Drewry Oviducal Entire and Jones, 1976). Each ovary Oviducal 5.1 contains one to six large fully yolked eggs and many (12-25) small (<0.6 mm diameter) im- mature ova. All of five females, collected variously from early April to the end of July, have two visible size classes of eggs in the ovaries. In addition, four of the five carry oviducal froglets in advanced developmental stages. The fat bodies.-Fat bodies in females carrying developing young are large flat lobes that fan out under the liver and over the oviducts. However, the fat bodies of the female that had fully yolked ovarian eggs but was not carrying young were short, thin finger-like structures. The oviducts.-The oviducts extend from the region of the heart, where they are held by connective tissue to the pericardium, to the cloaca. The anterior part of the duct, extending from the heart region to below the liver, is narrow and convoluted; the infundibular orifice is not expanded. The posterior part of the duct is a highly dilated thin-walled sac. The diameter of the dilated part of the duct of a female 20.5 mm snout-vent length (SV) carrying developing young is 8.8 mm. The straight-line length of the entire duct is 11.5 mm; the terminal 5 mm of the paired ducts are fused forming a common chamber to contain the developing young. The oviducts join the rectum and the urinarv ducts onlv 2.2 mm anterior to the vent. The cloaca1 region is therefore very small. In females carrying froglets, only the anteriormost 3 to 5 mm of the 12 mm duct is constricted and convoluted. In the female without froglets but with large ovarian eggs, some 10 mm of the duct is constricted and the left duct particularly is very convoluted. The ducts of all females are highly dilated posteriorly, and fused for some 5 mm, forming a common "uterus." LIVE-BEAR1NG ELEUTHERODACNLUS 123 GROSS MORPHOLOGY OF THE MALE REPRODUCTIVE TRACT The testes.-Testes are flat and round, 1.2-1.4 mm in diameter (three males examined). The testes of two of the three have punctate melanophores while those of the third appear unpigmented. The testes overlie the kidneys. The testes grossly appear to be comprised of several (10-15) lobes. The fat bodies.-The fat bodies of the specimen with unpigmented testes are large flat lobes; those of the specimens with pigmented testes are very reduced short, thin finger-like processes. I do not see any correlation of testis and fat body states with reproductive or nutritive condition. The ducts.-Thin archinephric ducts are visible leading to the cloaca from the testes. Ducts from the testes to the kidneys or the archinephric ducts cannot be discerned with the dissecting microscope, but probably extend from testis to kidney tubules since the testis overlies the kidney and is bound to it by connective tissue. EXTERNAL MORPHOLOGY OF THE FROGLETS All of the oviducal froglets examined are in advanced stages of development (Fig. 1). Fore and hind limb are well developed. The smallest froglet examined (4.6 mm SV) is the developmental equivalent of Lynn's (1942) stage IV; the largest is a 7.3 mm newly born free-living froglet, fully metamorphosed except for the presence of a tail rudiment and an as yet poorly developed tympanum, and the equivalent of Lynn's@ post-hatching stage V. The female carrying the oviducal young at the earliest stage of development had three fetuses in the left oviduct, two in the right. The two in the right duct were surrounded by membranes, probably the egg membranes. The others were free in the duct, closely appressed to the duct wall. Pairs of young in this female, and in the others dissected, were usually face-to-face and 'clasping' each other in a characteristic 'fetal' position with limbs drawn up to the body, back arched, and head % lowered.
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